天南星科Biarum的分类学修订

A TAXONOMIC REVISION OF BIARUM
Araceae

Peter C. Boyce

Summary. This paper contains a revision of the genus Biarum Schott. 21
species and 6 subspecies are recognised. All the taxa (except B. mendax Boyce)
are illustrated by line drawings, and most of them are shown as paintings or as
photographs in habitat or in cultivation. Distribution maps of all the species
are given and a key to all taxa is provided.

INTRODUCTION
Biarum comprises 21 species of dwarf tuberous-stemmed herbs
occurring in semi-arid and seasonally dry areas of southern Europe,
North Africa, the Near and Middle East. The centre of diversity
is the Middle East, where 75% of the species occur as endemics.
Biarum species have a strongly seasonal growth regime; the plants
begin growth in late summer or early autumn with the onset of winter
rains, continuing into late spring when they become dormant at the
start of summer heat and drought. The majority of species blossom in
autumn and early winter and this, together with the often striking ap-
pearance of the inflorescence, has resulted in the growing popularity of
Biarum species amongst alpine garden enthusiasts. In most species the
inflorescence emerges from the bare ground before the leaves (hyster-
anthous), but in some species the leaves are regularly synanthous, emerg-
ing at the same time as the inflorescence, and a few, such as Biarum
syriacum, are proteranthous, with the inflorescence emerging after the
leaves have developed. A taxonomic summary of the genus was given
in Aroideana (Boyce 2006). Further aspects of the horticultural popularity
of the genus have been covered by Mayo (1983) and Mathew (1987).

HISTORY OF THE GENUS
Until the publication of Schott’s articles in the Wiener Zeitschrift f ür
Kunst, Literatur, Theater und Mode (1829a, 1829b, 1829c, 1829d, 1829e,
1829f, 1829g; 1830a, 1830b, 1830c, 1830d, 1830e) and his aroid
account in Meletemata Botanica (1832), the genus Arum was ill-defined,
containing numerous species bearing only superficial similarity to
the type of the genus, A. maculatum L. Schott attempted to bring a
degree of homogeneity to Arum by segregating the anomalous taxa
in new genera. Schott (1832) proposed the genus Biarum to account

2 © The Board of Trustees of the Royal Botanic Gardens, Kew 2008.

for two species of Arum (sensu Linnaeus) with uniovulate ovaries and
a basal placenta, loosely arranged staminate ﬂowers with anthers
dehiscing by ventral longitudinal slits, scattered staminodes and
pistillodes and connate spathe tube margins. The two species included
were Biarum gramineum (Lam.) Schott and B. tenuifolium (L.) Schott.
Blume (1836) retained Schott’s two species and described an ad-
ditional three, Biarum bovei, B. homaïd and B. olivieri. Blume divided
Biarum into two sections, placing Schott’s species in “Sectio 1” (i.e.
sect. Biarum) and creating sect. Ischarum for the three new ones.
Blume distinguished sect. Ischarum by the lack of staminodes, anther
thecae dehiscing by apical pores and more-or-less elongated style.
By contrast, in sect. Biarum sensu Blume, both staminodes and pistil-
lodes are present, the thecae dehisce by longitudinal slits and the stigma
is sessile. Blume emphasized the different geographical distribution
of the sections as then known, with sect. Ischarum in the eastern Medi-
terranean region and sect. Biarum in the western Mediterranean.
Boissier (1853) added two new species, B. spruneri and B. alexan-
drinum, but did not review any of the previous treatments. He placed
B. spruneri in sect. Biarum (as sect. ‘Eubiarum’) with the note that
B. spruneri was apparently intermediate between sect. Biarum and
sect. Ischarum. Biarum alexandrinum was assigned to sect. Ischarum.
Schott and Kotschy (1854) raised Blume’s sect. Ischarum to generic
status with a single new species, I. eximium. No mention was made,
however, of the three species previously described by Blume for the
sect. Ischarum and Schott (1856) eventually made the necessary new
combinations in the genus Ischarum.
In his Synopsis Aroidearum, Schott (1856) published seven new com-
binations and two new species for Ischarum. The new combinations
included the two taxa described by Boissier (1853), Arum haenseleri
published by Willkomm (1847) and Biarum lehmannii Bunge (1851).
The last mentioned was later transferred to Eminium by Kuntze
(1891). The new species were I. kotschyi and I. dispar.
With the publication of Genera Aroidearum (1858) Schott proposed two
new genera, Cyllenium and Leptopetion, for Biarum spruneri and B. alexan-
drinum respectively. The differences between the genera concerned the
presence or not of a style, the means of thecae dehiscence, i.e. slits as
opposed to pores, and the shape and distribution of the pistillodes.
Engler (1879) adopted what was essentially Schott’s system ex-
cept that he reduced all Schott’s segregate genera to subgenera of
Biarum and dispensed with Leptopetion altogether, referring it to

© The Board of Trustees of the Royal Botanic Gardens, Kew 2008. 3

subgen. Ischarum. Engler also reduced many of Schott’s species to sub-
specific or varietal status or to synonymy.
Boissier (1882) also chose not to recognize Schott’s separate genera,
and in fact went a stage further than Engler (1879) in distinguishing
them at the rank of section rather than subgenus. Boissier followed
Engler in not accepting Leptopetion at any rank, also referring it to
section Ischarum.
Engler’s (1920) revision of Biarum for Das Pflanzenreich was the last
comprehensive treatment of the genus. Although little had changed
since his 1879 classification, he published one new species, B. straussii,
and a number of subspecific taxa.
Since the Pflanzenreich account several floristic examinations of
Biarum have been undertaken by various authors (e.g. Riedl 1963
& 1985, Mouterde 1966, Talavera 1976, Mill 1984 and Koach &
Feinbrun 1986). Riedl (1980b) published a preliminary summary
of the genus together with a key to the species. However, none of
these accounts attempts a comprehensive treatment and new dis-
coveries and interpretations during the last fifty years have made
this revision necessary.

TYPIFICATION
During the course of this revision it has been necessary to lectotypify
a number of names. The majority of these are names published by
Schott for which the types in Vienna (W) were destroyed during the
closing stages of the Second World War. For lectotypes I have followed
the advice of Riedl & Riedl-Dorn (1988) in selecting illustrations
that were commissioned by Schott and prepared from living and
herbarium specimens. These are known collectively as the Icones &
Reliquiae Aroidearum. It has also been necessary to lectotypify names based
on types destroyed in Berlin. In these cases it has not been possible
to trace any authentic material, but in some instances drawings of
the types exist and have been chosen to serve as the lectotype.

FUTURE RESEARCH
Unlike the genus Arum, practically nothing has been done in
connection with studying the pollinators of Biarum. The major
reason is undoubtedly the inconvenient flowering period (from the
botanical standpoint) of most of the genus, which, coupled with
the fleetingness of the inflorescences and the often widely scattered
individual plants, makes them difficult to study effectively. Koach


Biarum carduchorum showing pollination by Staphylinid beetles. Photograph by Peter Boyce.

(1986) in his study of the Israeli Araceae makes no comment as
to pollinator type in Biarum. B. carduchorum (in Turkey at least)
attracts Staphylinid beetles in some considerable numbers (pers. obs.),
although it is unclear whether it is these, or the numerous Diptera
also attracted by the foetid odour, that facilitate pollination. A similar
situation exists with Greek populations of B. tenuifolium and B. spruneri
and I strongly suspect that much the same is true for all the foetid
species. Certainly many of the herbarium specimens I have seen
during the course of this revision have had beetles, small ﬂies and
midges in the lower spathe. The situation with the sweet-smelling
B. davisii is quite different. Observations of both the typical subspe-
cies in Crete and subsp. marmarisense on the island of Simi in the


eastern Aegean have shown that bees and queen wasps are at-
tracted by the sweet lilac-like perfume emitted by the inflorescences
(pers. obs).
Some work has been done with the pollen of Biarum and so far
the results suggest that palynology may not prove to be of significant
use. If so, this is similar to the situation in Arum where there is
depressingly little variation in the types of pollen-exine sculpture
between species. Talavera (1976) studied four species in Spain and
Portugal and showed them to have similar spinose pollen and Grayum
(1986) found that there is little to separate the pollen between
Ischarum and Biarum. Pollen of the morphologically distinct B. davisii
is also quite similar to other species. However, B. ditschianum Bogner
& Boyce (1989) has smooth pollen, suggesting that a complete survey
of the pollen may provide useful data.
A more promising field of research is cytology. Although less
than a third of the species have been studied to date the counts
obtained (e.g. Talavera 1976) certainly support the separation of
some of the more controversial taxa. In particular the problematic
Afro-Iberian species are cytologically different from the eastern
Mediterranean species that they are often merged with. At the
present time research is being undertaken by Athanasiou in Patras
University, Greece, into the cytology of the Greek taxa and already
results have helped with the delimitation of some taxa and data
has been used in this revision. To date the following counts have
been found for the genus: 2n = 16, 20, 22, 24, 26, 32, 36, 74,
c. 96, 98.
All specimens cited have been seen by the author unless otherwise
stated.

TAXONOMICALLY IMPORTANT CHARACTERS
The following characters are of importance in Biarum taxonomy at
subgeneric level:-
1. Staminate flower structure, i.e. beaked or globose-oblong.
2. Dehiscence type, i.e. ventral longitudinal slit as opposed to
apical pore.
3. Presence or absence of staminodes.
4. Shape of pistillodes, e.g. filiform or hooked.
5. Seed shape and size, e.g. ovoid or globose, more or less than
3.5 mm diam. or more or less than 5.5 mm.


The following characters are of importance in Biarum taxonomy
at sectional level:-
1. Spathe tube shape and size in proportion to the spathe
limb.
2. Spadix appendix shape, i.e. fusiform to slender filamentous
and presence or not of additional epidermal structures.
3. Inflorescence odour, e.g. foetid-smelling, sweet-smelling or
odourless.
The following characters are of importance in Biarum taxonomy
at specific level:-
1. Degree of spathe tube margin connation.
2. Distribution of pistillodes.
3. Staminate-pistillate flower zone interstice length.

SUPRAGENERIC CLASSIFICATION
Biarum is interesting because, despite its Euro-Mediterranean distribu-
tion, it is not closely allied to other Araceae occurring in the area, dis-
playing instead similarities to the Asiatic genera Theriophonum Blume,
Typhonium Schott and Sauromatum Schott. It is my belief that Biarum
has derived from a separate ancestral stock to Arum and Eminium, the
genera most closely associated with Biarum in its natural range.
In attempting to understand the relationships between Biarum
and rest of the Arinae it is logical to investigate first the other Medi-
terranean genera belonging to the subtribe. The difficulty is that
on the basis of general morphology none of these genera are
apparently closely related to Biarum.
The simple leaves of Biarum have no parallel in the Mediterranean
Arinae. One genus, Eminium (Blume) Schott, has two Central Asian spe-
cies with lanceolate undivided leaves, but all the Mediterranean Eminium
species have helicoid-pedatifid leaves. Arum euxinum Mill, a species from
north western Turkey, has leaves with poorly developed posterior lobes
and occasionally produces leaves that are more-or-less unlobed. I would
not suggest, however, that there is a link between A. euxinum and Biarum
since the leaf shape is the only common character; the inflorescences
and infructescences of A. euxinum are otherwise typical for Arum.
If inflorescences are compared, the closest genus to Biarum would
appear to be Eminium which with its ground-level or partially buried
vernal inflorescences is superficially similar to some species of Biarum,


for example the spring flowering B. syriacum (Spreng.) H.Riedl
and B. straussii Engler. Aside from these phenological similarities,
both Eminium and Biarum have uniovulate ovaries with a basal
placenta. Well developed pistillodes are present in most species of
Biarum and all species of Eminium and most taxa in both genera
have a strongly foetid spadix appendix. In addition, the shape of
the pistillodes in Eminium is very similar to that in Biarum subgenus
Biarum. With regard to fruiting, both genera have subterranean to
partially emergent infructescences consisting of silver-lilac to off-white
berries and seeds with strongly developed elaiosomes. Both genera
are adapted to seasonally arid conditions. Biarum and Eminium differ
primarily in the staminate flowers, which, in Eminium, show none
of the diversity present in Biarum and in fact are virtually indistin-
guishable from the staminate flowers present in Arum. There is also
some disparity with regard to the pistillate flowers; the oblong ovaries
with sessile stigmas found in subgen. Biarum and some taxa in subgen.
Ischarum are not dissimilar to the ovaries in Eminium, but the bottle-
shaped ovaries with a stigma borne on a long style that are typical
of the majority of Biarum are quite unlike anything known in
Eminium, or in the rest of the Mediterranean Arinae. It is also notable
that the hemispherical arrangement of the pistillate flowers in Biarum
is quite different to the oblong cluster of ovaries found in Eminium
and in the other genera of the Arinae that occur in the area.
Arum, with parietal placentation, a highly evolved pollination sys-
tem, hastate to sagittate leaves and displayed infructescences is, in my
opinion, morphologically too far removed from Biarum to be anything
but a distant relative. Biarum, for all its floral diversity, has a fairly
simple pit-fall inflorescence with none of the specialization present
in Arum (see Boyce 1991). Furthermore, in the Arinae parietal placen-
tation is known only in Arum, all of the other genera have basal pla-
centas (e.g. Biarum, Eminium, Sauromatum Schott, Typhonium Schott) or
basal and apical placentas (e.g. Dracunculus Miller, Helicodiceros Schott,
Theriophonum Blume). At this juncture it is worth noting that, despite
the anomalous autumn flowering strategy and the anthers dehiscing
by apical pores, it seems unlikely that A. pictum represents a close link
between the genus Biarum and the remainder of Arum. These features
aside, A. pictum is, morphologically, a fairly typical Arum.
The other Mediterranean representatives of the Arinae, Dracunculus
and Helicodiceros, are probably the least tenable as regards a close af-
finity with Biarum. Both Dracunculus and Helicodiceros show considerable


morphological dissimilarities to Biarum which include a well developed
pseudostem, strongly divided leaves, apical and basal placentation,
few-ovulate ovaries and large, displayed, infructescences. The sta-
minate flowers in Dracunculus and Helicodiceros are virtually indistin-
guishable from those in Arum and Eminium, as too are the pistillate
flowers.
Engler’s opinion underwent an important change concerning the
position of Biarum relative to the other genera of the subtribe Arinae.
Until the Pflanzenreich treatment, he had placed Biarum before Arum,
suggesting that he regarded Biarum as less specialized than Arum.
In the Pflanzenreich, however he placed the genus at the end of the
Arinae after Eminium.
It is my belief that the closest links with Biarum lie with Typhonium,
Theriophonum and Sauromatum. Vegetatively there are few similarities
between Biarum and these three genera, although the simple leaves
of Biarum have parallels in the Indian and Sri Lankan genus
Theriophonum (T. sivaganganum (Ramam. & Seb.) Bogner and in the
juvenile leaves T. fischeri Sivadasan) and in two Australian Typhonium
species (T. liliifolium F. Muell. and T. alismifolium F. Muell.). Repro-
ductively, however, Sauromatum, Typhonium and Theriophonum share a
number of common features with Biarum such that it is difficult to
determine to which Biarum is closest. The staminate flower structure
of Biarum is variable. The majority of species possess two thecae
joined by an inconspicuous connective, similar to Sauromatum and
many Typhonium species, although subgen. Biarum and subgen.
Cyllenium have elongated connectives of a type found in some Typhonium
species, e.g. T. flagelliforme (Lodd.) Blume and in Theriophonum minutum
(Willd.) Baillon, a species widespread in India and Sri Lanka.
In Biarum, beaked connectives are associated with thecae dehiscing
by longitudnal slits while in Typhonium and Theriophonum they are
associated with dehiscence via apical pores.
Looking to pistillode structure, the odd, club-shaped pistillodes
found in Sauromatum and some species of Typhonium (e.g. T. flagelliforme)
are unknown in Biarum, where the pistillodes are usually filiform
(in subgen. Ischarum) or thickened and hooked (in subgen. Biarum).
This latter type of pistillode is also found in Theriophonum, specifically
in T. sivaganganum.
The pistillate flowers of Biarum show some similarity to those in
Theriophonum as well as to some species of Typhonium. On the basis
of placentation and ovule number Typhonium would appear to be


closest to Biarum, in possessing a basal placenta with one or two
ovules. Sauromatum has similar placentation but generally has between
two and four ovules per ovary. Theriophonum has apical and basal
placentation with up to six ovules per locule. One feature not found
in any of the genera under discussion, except Biarum, is a distinctive
style. The stigmas in Sauromatum, Typhonium and Theriophonum are
always sessile. It should be noted, however, that this character is
not found in all Biarum species and that species with sessile stigmas
are found in both subgenera.
The connation of the spathe-tube margins that features prominently
in Biarum is found only in Sauromatum. To date no species unequivo-
cally attributable to Typhonium or Theriophonum has been found to
possess this character. The odd, hairy-leaved Typhonium hirsutum
(S.Y. Hu) J. Murata & Mayo, which is notable in having partially
connate spathe-tube margins, will require more study before its posi-
tion can be properly evaluated. Interesting also is Sauromatum brevipes
(Hook.f.) N.E. Br. a remarkable, little-collected species from Sikkim.
In leaf S. brevipes is similar to a dwarf plant of S. venosum (Ait.)
Schott but in ﬂower the similarity to Biarum davisii Turrill, both in
shape and colouration, is quite remarkable.
CULTIVATION. Biarum species can be divided into three groups for
the purpose of cultivation. Some species, notably Biarum tenuifolium
and some of its allies, are easy to grow and in sheltered positions
in milder parts of Europe are hardy when planted in the open.
However, the majority of species require some winter protection,
ideally frost free, and are best grown in pots in a cool greenhouse
or planted directly into a sheltered cold frame. Lastly there are a
few species that tax the grower and require cosseting in a cool
greenhouse; Biarum olivieri is typical of this third group.
Biarum tenuifolium is one of three species, the others being B. arundanum
and B. rhopalospadix, which can be successfully grown outdoors in
favourable areas. The tubers require quite deep planting, 5–7 cm
down, and a rich well drained soil. If the soil tends towards clay
then the addition of sharp sand and grit, particularly as a cushion
beneath the tuber, is beneﬁcial. In warm areas the tubers can be
planted in open places, in a rockery for example, but in less ideal
situations a planting site at the base of a west or south facing wall
will provide the best chance of success. When happy, B. tenuifolium
in particular will spread freely both by offsets and from seed. Not
all subspecies of B. tenuifolium are equally successful out doors. Subsp.


Biarum marmarisense cultivated at the Royal Botanic Gardens, Kew. Photograph Richard Wilford.


tenuifolium and subsp. abbreviatum are best, while subsp. zelebori and
particularly subsp. idomenaeum require extra care in siting and vigilance
to ensure good growth. B. rhopalospadix grows well in conditions
similar to those which suit B. tenuifolium.
In particularly sheltered places with a Mediterranean-like climate
other species that are worth experimenting with outdoors include
B. crispulum, B. davisii, B. fraasianum, B. kotschyi and B. marmarisense.
All the above, together with most of the other species, will do
very well in pots in a frost-free greenhouse. Most do best in rather
deep pots with the tuber planted quite deeply (3–4 cm) in an open
rich loam-based soil. If sufficient tubers are available most species
seem to be better planted several to a large, deep clay pot. Gener-
ally plants grown in solitary confinement, one to a pot, seem less
vigorous. Annual repotting is advantageous and also permits an
inspection of the tubers for signs of rotting. Potting is best carried
out in summer (before August). The later it is left the more chance
there is of root damage occurring. A good mix is: 45% loam based
compost, 40% humus, 15% potting grit (all percentages by volume).
The mix should be damp at the time of potting. The pots should
be filled to within 2 cm of the rim and then topped off with grit.
This will allow moisture to drain away from the ‘neck’ of the plants.
After potting place the pots somewhere cool and do not water until
growth becomes visible.
Once growth has begun, water sparingly until flowering is over
or growth is beginning to accelerate. Too much water early on can
result in root loss. It is good to encourage the plants to grow as
vigorously as possible before the start of winter. This will ensure
that the plants remain stout and that a good sized tuber is produced
for the next season. Watering should be continued and the plants
not allowed to become dry at the roots. A feed at every watering
with a fertilizer designed for tomatoes will encourage robust growth.
Keep the greenhouse or frame well ventilated at all times. This will
discourage damp-related problems such as Botrytis.
Plants grown at Bonn, Munich, Kew and in a number of private
collections in Germany and the UK succeed best in a relatively
narrow but deep clay pot – i.e. a ‘long-tom’ – in a compost of
equal parts sterilized loam, humus and sharp grit. Under glass
the plants should be given as much light as possible during the
winter to prevent leaf etiolation. Ventilation must be given on all
but the coldest days, and water applied regularly but carefully, since


moisture remaining lodged in the leaf sheaths can result in
rotting. The plant should continue growing throughout the winter.
Towards early spring growth will slow down and by mid-June
the leaves will turn yellow and wither. At this time watering should
be more or less stopped. The compost should be kept just moist
by plunging the pot to the rim in sand and keeping the sand damp.
In some species the inflorescence will emerge soon after the foliage
dies, and in these, once flowering is over, the compost should be
allowed to dry out and the plant rested. Care should be taken not
to allow the pot to become too hot, as this can cause desiccation
of the tuber. Propagation is best effected by seed, although some
species produce offsets which can be removed during repotting
and grown on.

TAXONOMIC TREATMENT
Biarum Schott nom. cons. in Schott & Endlicher, Melet. Bot. 17 (1832) &
Syn. Aroid. 6 (1856) & Gen. Aroid. t.7 (1858) & Prodr. Syst. Aroid. 60–64
(1860); Pfeiffer, Nomen. Bot. 1(1): 403–404 (1873); Engler in A. & C. DC.,
Monog. Phanerog. 2: 571–580 (1879) & in Engler & Prantl., Die Natürl.
Pflanzenfam. 149 (1889) & in Engler, Pflanzenr. 73(IV.23F): 132–143
(1920); Riedl in Aroideana 3(1): 24–31 (1980); Mayo, Bogner & Boyce,
The Genera of Araceae, 266–268, pl.96 (1997). Type: B. tenuifolium (L.)
Schott.
Homaïd Adans., Fam. Pl. 2:470 (1763) in syn. nom. rejic.; Pfeiffer, Nomen. Bot.
1(2): 1658 (1874). Type: H. tenuifolium (L.) Adans.
Ischarum Schott & Kotschy, Oesterr. Bot. Wochen. 4: 81 (1854); Schott, Syn.
Aroid. 6–8 (1856) & Gen. Aroid. t.10 (1858) & Prod. Syst. Aroid. 65–70
(1860); Pfeiffer, Nomen. Bot. 1(2): 1764 (1874). Type: I. bovei (Blume) Schott
(See Nicolson 1967).
Leptopetion Schott, Gen. Aroid. t.8 (1858) & Prodr. Syst. Aroid. 64 (1860).
Type: L. alexandrinum (Boiss.) Schott.
Cyllenium Schott, Gen. Aroid. t.9 (1858) & Prodr. Syst. Aroid. 64–65 (1860).
Type: C. spruneri (Boiss.) Schott.
Stenurus Salis., Gen. Pl. 5 (1866). Type: S. tenuifolium (L.) Salis.
Homaida Adans. emend O. Kuntze, Rev. Gen. Pl. 2: 742 (1891). Type:
H. tenuifolium (L.) Adans. emend O. Kuntze.

DESCRIPTION. Dwarf tuberous stemmed herbs. Tuber dorsoventrally
compressed-discoid to ± globose, encased basally by remains of the previous
season’s tuber, growth point central, peripheral adventitious buds none to
many, usually giving rise to independent tubers, tuber apex coated with
moderate to copious amounts of waxy farina and bearing the remains of
the previous season’s cataphylls; roots simple, emitted in a ring around the
growth point, contractile or feeding; contractile roots fusiform, usually thick,


feeding roots slender, cylindrical. Foliage hysteranthous, occasionally synan-
thous, rarely proteranthous, petiolate; petiole sometimes ± subterranean,
leaves erect or, rarely, reflexed; petioles enclosed proximally by several cata-
phylls, the inner papery and the outer fibrous, these often emerging above
ground and encasing the basal part of the aerial shoot, petioles terete proxi-
mally, canaliculate and expanded into a membranous wing distally, petioles
enclosing one another, forming a weakly defined aerial pseudostem in some
species; leaf lamina linear, lanceolate, elliptic-oblong or spathulate, decurrent,
± cuneate, rounded or ± truncate apically, acute to obtuse basally, margins
smooth to undulate, rarely crispulate, mid-, light, bright or deep green, rarely
with green or black-purple bullae or silver-grey spotting. Inflorescence ± sessile
on the tuber or borne on a short to rather long, subterranean peduncle,
rarely peduncle emerging above ground, smelling foetid or sweet. Spathe
divisible into a tubular lower portion (spathe tube) and a ± expanded upper
portion (spathe limb); spathe tube narrow to inflated, sometimes greatly so,
constricted distally or not, emergent to partially buried, rarely entirely sub-
terranean, margins partially to fully connate, sometimes convolute distally,
exterior deep purple to dirty green or greenish purple above, ± white below,
interior off-white below, purple above, or wholly purple or off-white with
purple staining at the base around the pistillate flowers; spathe limb large to
small, rarely ± absent, linear, lanceolate or elliptic, erect, reflexed or deflexed,
flat to recurved and coiled or strongly involute; exterior dirty green, more
rarely mid-green, dirty white, dull yellow or pinkish brown sometimes with
purple spotting and staining; interior deep purple-brown, yellow or pale
greenish, sometimes purple with a green apex, apex subacute, acute to
acuminate, margins smooth to undulate, rarely crispulate. Spadix shorter than,
equalling or exceeding the spathe limb, divided into distinct zones; a sterile
terminal appendix, an zone of staminodes (subgenus Biarum), a fertile male
zone, a stamino-pistillate interstice, a further zone of staminodes (sometimes
absent) and a fertile female zone; terminal appendix sessile to stipitate, cylindri-
cal to fusiform, erect to flexuous, apex acute to rounded, tapering, base rarely
rounded or sub-truncate, smooth, very rarely furnished with filamentous
processes proximally, deep purple, brown-red or brown, occasionally greenish,
rarely dirty yellow. Flowers: supra staminal staminodes present only in subgen.
Biarum, arranged in few to several irregular whorls, simple or 1–2-branched,
hooked, peg-like or filamentous, partially expanded proximally, glossy, cream;
staminate flowers arranged in a cylindrical, globose or oblong zone, each com-
prising two anthers, connective short to ± absent, rarely prolonged into a ±
prominent beak, thecae dehiscing by coalesced or separate apical pores
(subgen. Ischarum) or ventral longitudinal slits (subgen. Biarum), cream to
purple; pollen free or in strands, exine spinose or smooth, interflora staminodes
usually confined to the base of the stamino-pistillate interstice, more rarely
arranged evenly over the whole area or absent, slender-filiform to hooked,
simple to 1–3-branched, recurved, decurved or tangled, cream, off-white or
purple; pistillate flowers arranged in a hemispherical cluster at the base of the
spadix, ovary oblong, sub-globose or bottle-shaped, off-white to purple,
uniovulate with a basal placenta, ovules orthotropous, style slender to rather


stout or absent, stigma capitate, pale greyish or purple. Infructescence subter-
ranean or partially emergent, globose berries many, globose to pyriform,
white to lilac- or purple-stained, stigmatic remnents slightly prominent or
not; seed ovoid to globose, large to small with a large elaiosome at the hilum,
testa leathery, ± smooth to reticulate, pale to dark brown, endosperm copious,
embryo straight; eophyll lanceolate to spathulate.
Twenty-one species of dwarf tuberous-stemmed strongly seasonal herbs
occurring in semi-arid and seasonally dry areas of southern Europe, North
Africa, the Near and Middle East.
Subgenus Biarum
Species:
1a. tenuifolium subsp. tenuifolium
1b. tenuifolium subsp. arundanum
1c. tenuifolium subsp. galianii
1d. tenuifolium subsp. zelebori
1e. tenuifolium subsp. abbreviatum
1f. tenuifolium subsp. idomenaeum
2. rhopalospadix
Subgenus Ischarum
3. aleppicum
4. angustatum
5. carduchorum
6. eximium
7. bovei
8. crispulum
9. dispar
10. olivieri
11. straussii
12. syriacum
13. carratracense
14. kotschyi
15. fraasianum
16. pyrami
17. mendax
18. auraniticum
19. ditschianum
20. davisii
21. marmarisense

KEY TO SPECIES
1. Staminodes occurring above and below the staminate ﬂower zone ............. 2
Staminodes absent above the staminate ﬂower zone; present or absent
below ........................................................................................................ 7
2. Staminodes hooked ................................................................................. 3


Staminodes not hooked ........................................................................... 6
3. Staminodes simple, very rarely branched .............................................. 4
Staminodes always 2–3 branched ........................................................... 5
4. Leaf lamina 15–25 cm ´ 11–15 mm; spadix appendix 10–41 cm ´ 2–3
mm; leaves in mature individuals oblong-lanceolate early in the season,
linear-lanceolate later in the season .........................................................
................................................. 1a. tenuifolium subsp. tenuifolium
Leaf lamina 20–40 cm ´ 16–21 mm; spadix appendix 10–12 ´ 4–9 mm;
leaves in mature individuals always oblong-lanceolate. ...........................
......................................................... 1d. tenuifolium subsp. zelebori
5. Leaf lamina oblanceolate to linear-lanceolate; staminodes 2-branched,
arranged in c. 8 regular whorls. Plants of heavy terra rossa soils .........
................................................. 1b. tenuifolium subsp. arundanum
Leaf lamina linear; staminodes 2–3-branched, arranged in c. 7 irregular
whorls. Plants of loose sandy soils ............................................................
...........................................................1c. tenuifolium subsp. galianii
6. Staminodes peg-like; leaf lamina spathulate, erect, margins gently undu-
late .........................................1e. tenuifolium subsp. abbreviatum
Staminodes filamentous; leaf lamina linear-oblong, usually adpressed to
the ground, margins strongly undulate-crispulate ....................................
................................................ 1f. tenuifolium subsp. idomenaeum
7. Staminodes hooked, stiff, thickened. Thecae dehiscing by ventral longi-
tudinal slits, connective rostrate ................................ 2. rhopalospadix
Staminodes filiform, flexuous, or absent, never hooked and thickened.
Thecae dehiscing by apical pores, connective barely prominent or flush
with the anther surface ........................................................................... 8
8. Spadix appendix massively thickened with reflexed basal ‘hairs’; spathe
limb much reduced, recurved ................................. 19. ditschianum
Spadix appendix with no ornamentation; spathe limb well developed or,
if reduced, then erect, cucullate ............................................................. 9
9. Spathe limb usually much shorter than the spathe tube, margins recurved,
spathe tube enclosing much of the spadix ............................................... 10
Spathe limb exceeding the spathe tube in length, flat or with the margins
inrolled, spathe tube enclosing the base of the spadix only ............... 11
10. Spathe 5–6 cm long, spadix appendix 3–3.5 cm ´ 2 mm ...20. davisii
Spathe 7–8 cm long, spadix appendix 3.5–5 cm ´ 0.5 mm ...................
.................................................................................. 21. marmarisense
11. Spathe tube not inflated, the sides ± parallel ...................................... 12
Spathe tube inflated, the sides ± gibbous ............................................ 14
12. Spathe tube margins free ± to the base; stigmas not borne on a stipe ..
........................................................................................... 3. aleppicum
Spathe tube margins free for a quarter of their length; stigmas borne on
a short to moderately long stipe ........................................................... 13
13. Staminodes directed downwards; leaves narrowly lanceolate-elliptic .......
.........................................................................................4. angustatum
Staminodes directed upwards; leaves elliptic to broadly oblong-elliptic
....................................................................................... 5. carduchorum


14. Spathe tube distinctly wider than the spathe limb, the margins joined for
their entire length; spathe limb appearing linear due to inrolled margins;
spadix appendix filiform ....................................................... 10. olivieri
Spathe tube as wide as or narrower than the spathe limb, the margins
free for at least a quarter of their length ............................................ 15
15. Foliage proteranthous ............................................................................ 16
Foliage hysteranthous or synanthous .................................................... 17
16. Leaf lamina ovate-elliptic, oblong or linear; staminodes restricted to the
bottom quarter to third of the interstice .......................... 11. straussii
Leaf lamina in mature plants linear to linear-elliptic; staminodes distributed
over the basal half of the interstice ........................................ 12. syriacum
17. Staminodes distributed evenly over the interstice separating the staminate
and pistillate flower zones ................................................... 6. eximium
Staminodes arranged mostly above the pistillate flower zone, intersticial
staminodes adjacent to the staminate flower zone much reduced or ves-
tigial, or staminodes absent ................................................................... 18
18. Spathe limb interior greenish white, spadix appendix yellow .................
......................................................................................18. auraniticum
Spathe limb interior deep purple-brown; spadix appendix similarly
coloured.................................................................................................. 19
19. Spathe tube margins free for quarter to half their length .......................... 20
Spathe tube margins free for three quarters their length ....................... 21
20. Spathe tube margins free for half their length, staminodes few, SW
Spain ..........................................................................13. carratracense
Spathe tube margins free for a quarter of their length, S Greece .........
........................................................................................ 15. fraasianum
21. Spadix appendix c. 2–4 mm in diam., slender-cylindric, to slender-fusiform,
appearing ± consistent diameter .......................................................... 22
Spadix appendix more than 4 mm in diam., fusiform, widest below the
middle .................................................................................................... 24
22. Spathe tube globose; interstice twice as long as the staminate flower zone;
staminodes very few or absent ................................................ 9. dispar
Spathe tube oblong; interstice equalling the staminate flower zone; sta-
minodes many ........................................................................................ 23
23. Foliage hysteranthous; spathe limb lanceolate, margins smooth .............
..................................................................................................... 7. bovei
Foliage synanthous; spathe limb linear-lanceolate, margins crispulate ...
.............................................................................................8. crispulum
24. Spathe tube oblong, spathe limb narrowly lanceolate, acute, seldom ex-
ceeding 10 cm .................................................................... 14. kotschyi
Spathe tube globose, spathe limb lanceolate, acuminate, exceeding 12 cm
................................................................................................................. 25
25. Spathe tube margins fully fused; spathe limb remaining erect during flow-
ering ..................................................................................... 17. mendax
Spathe tube margins for ¼ of their length ; spathe limb reflexing and
curling during flowering ....................................................... 16. pyrami


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